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AUSTRALIAN MUSEUMSCIENTIFIC PUBLICATIONS

Australian Museum science is freely accessible online at

www.aust ra l ianmuseum.net .au/publ icat ions /

6 College Street, Sydney NSW 2010, Austral ia

nature culture discover

Hand, Suzanne, Lyndall Dawson and M. Augee, 1988. Macroderma koppa, a new Tertiary species of false vampire bat (Microchiroptera: Megadermatidae) from Wellington Caves, New South Wales. Records of the Australian Museum 40(6): 343–351. [31 December 1988].

doi:10.3853/j.0067-1975.40.1988.160

ISSN 0067-1975

Published by the Australian Museum, Sydney

Records of the Australian Museum (1988) Vol. 40: 343-351. ISSN 0067 1975 343

Macroderma koppa, a new Tertiary species of false vampire bat (Microchiroptera: Megadermatidae)

from Wellington Caves, New South Wales

SUZANNE HAND, LYNDALL DAWSON & MICHAEL AUGEE

School of Zoology, University of New South Wales, Kensington, NSW 2033, Australia

ABSTRACT. A new late Tertiary false vampire bat, Macroderma koppa, is described from the Big Sink doline of Wellington Caves, eastern central New South Wales. The new species appears to be the sister-group of the living Australian Ghost Bat, Macroderma gigas. Morphological features that distinguish the new species from M. gigas appear to be plesiomorphies shared with most other megadermatids.

HAND, S., L. DAWSON & M. AUGEE, 1988. Macroderma koppa, a new Tertiary species offalse vampire bat (Microchiroptera: Megadermatidae) from Wellington Caves, New South Wales. Records of the Australian Museum 40( 6): 343-351.

Until recently, all Australian fossil megadermatids had been identified as conspecific with (or very close to) the only living Australian megadermatid, the Ghost Bat M aeroderma gigas. These fossils come from late Pleistocene and Holocene deposits in south-western Western Australia, the Flinders Range in southern central South Australia, north-eastern Queensland and north-eastern and eastern central New South Wales (see Molnar et al., 1984; Fig. 1). Recently, the remains of Miocene species of the Australian endemic genus Macroderma have been recovered from limestone sediments on Riversleigh Station, north-western Queensland (Hand, 1984, 1985, 1987). In this paper a new late Tertiary species of Maeroderma from New South Wales is described.

Fossil specimens described here were collected from breccia in the Big Sink, one of a complex of cave deposits occuring in Wellington Caves, New South Wales. These caves are formed in limestones of the Devonian Garra Formation which outcrops in low hills 1 km east of the Bell River approximately 7 km south of the town of Wellington (32 0 35'S, 148 0 59'E). They comprise at least five natural caves which have been expanded and greatly disturbed by fossil collection and phosphate mining over a period of 150 years. The history of fossil collection and mining from this locality has been described in detail by Dawson (1985). Cave nomenclature used here follows Dawson (1985, fig. 2).

Recent geological studies of the Wellington cave fills (Frank, 1971; Osborne, 1983) have indicated their considerable stratigraphic complexity. Osborne (1983) has hypothesised a depositional sequence for various stratigraphic units in the Wellington Caves system but the absolute ages of the units have not been determined. Osborne (1983), following Frank (1971), Francis (1973) and others, suggests that the oldest deposits within the caves could have been laid down in the Miocene. He cites faunal evidence suggesting that deposition of bone breccia occurred throughout the Pleistocene at least (Osborne, 1983).

Stratigraphically controlled excavations in the caves were made by the School of Zoology of the University of New South Wales in 1982-1987. Some preliminary results of this collection have been reported by Dawson (1985). Material described here was collected in October 1982 by M. Archer and students of the School of Zoology, University of New South Wales, from the Big Sink doline as part of a pilot study.

The species recovered from the Big Sink doline include : the macropodine Protemnodon sp. cf. P. devisi (the most abundant species); a plesiomorphic macropodine allied to Wallabia spp.; Maeroderma koppa n.sp.; Thylacinus sp.; Thylaeoleo crassidentatus; Petauroides stirtoni (a pseudocheirid otherwise known from the Hamilton local fauna of Victoria); several small dasyurids which are either

344 Records of the Australian Museum 40

I ...

I ... Ai

I ... I ... ...

r----L-, I

V Fig.I. Map of Australia showing the distribution of Macroderma (solid triangles, extant populations of M. gigas; open triangles, fossil and subfossil remains). Open circles on map represent localities discussed in the text: 1, Cliefden Caves; 2, Wellington Caves; 3, Yessabah Caves; and, 4, Ashford Cave. (After Molnar et al., 1984.)

new species or species otherwise poorly known from late Tertiary deposits (e.g. Antechinus sp. comparable to that described from the Floraville local fauna, Queensland by Archer 1982); a small bandicoot which probably represents a new Tertiary genus; a burramyid; and a new pseudomyine rodent. The non-mammalian fauna includes a species of Tiliqua, a scincid. The fauna is believed to be early to middle Pliocene in age (see Discussion) and is being described by L. Dawson (in prep.).

Institutional abbreviations used here are as follows: AM F - Australian Museum Fossil Collection; AM M - Australian Museum Mammal Collection (see Fig. 4); BMNH - British Musem (Natural History); UNSW WC - University of New South Wales, Zoology Museum, Wellington Caves Fossil Collection; WAM - Western Australian Museum (see Fig. 3).

CHIROPTERA Blumenbach, 1779

MICROCHIROPTERA Dobson, 1875

RHINOLOPHOIDEA Gray, 1825

MEGADERMATIDAEAllen, 1864

Macroderma Miller, 1907

Macroderma koppa n.sp. Type material. HOLOTYPE (Fig. 2): the holotype (AM

F69070) is a partiaf skull with associated dentaries. It

contains a complete dentition, i.e. 11 2' Cll' p424, Ml.3 l_3· PARA TYPES: a number of specimens have been selected as paratypes because each serves to illustrate features not represented in the holotype. The paratypes are : UNSW WC171, a skull fragment preserving the cheek, palate and part of the nasal regions; UNSW WC153, fragment with MI-3 and the maxillary base of the zygomatic arch; UNSW WC159 fragment preserving the back ofthe palate and pc M3; UNSW WC158, fragment with P4_M3; UNSW WC170, maxillary fragment preserving the cheek region and P4_M2; UNSW WC173, 174 and 175, fragments of right Cls; UNSW WC176, fragment of left Cl; UNSW WC156, associated dentaries with I l-M3; UNSW WC173, dentary preserving P 4-M3' alveoli for Cl and P 2 and base of ascending ramus; UNSW WC152 dentary fragment preserving MI_3 and base of ascending ramus; UNSW WC165, MI_3; UNSW WCI77, a right Cl; UNSW WC178, a fragment of a left Cl; and UNSW WC179, a juvenile dentary without teeth. Other referred specimens are also lodged in the Zoology Museum at the University of New South Wales.

Diagnosis. The new species is referred to the megadermatid genus Macroderma which currently includes two species: the living Australian Ghost Bat Macroderma gigas and the extinct M. godthelpi of the Miocene of north-western Queensland (Hand, 1985). Species of Macroderma may be distinguished from other megadermatids (of the genera Lavia, Cardioderma, M egaderma, Lyroderma and the extinct monotypic Necromantis) by, among other features, their large size, enlarged heels of P4_M2,

Hand, Dawson & Augee: Macroderma koppa 345

MM • • • • • Fig.2. Macroderma koppa n.sp., AM F69070 (holotype), partial skull with associated dentaries and complete dentition. a-al, stereopairs, buccal-oblique view of mandibles with 11,12, Cl' P2, P4, MI, M2, M3; b, buccal view, right maxillary fragment; and, c-c l , stereopairs, occlusal-oblique view, right maxillary fragment with Cl, P4, MI, M2 and M3. Photos by R. Oldfield.

346 Records of the Australian Museum 40

more prominent paracone of Cl and more robust buccal cingula of the lower molars (Hand, 1985).

The fossil species differs from the living M acroderma gigas in its retention of two infraorbital foramina, its poorly-developed nose-shield, its larger incisors and premolars, its less laterally compressed P , its lower-crowned and more posteriorly recurved pj, and in its proportionately longer anterior tooth row.

Macroderma koppa differs from the Australian Miocene species M. godthelpi in its much larger size, relatively larger posterior infraorbital foramina, loss of p2, well-developed p4 heel, its higher crowns and its more reduced pre-entocristids.

Type locality, lithology and age. The described material was collected from the southern wall of the Big Sink doline at Wellington Caves, NSW. Breccia from this position was identified by Osborne (personal communication, 1982) as part of the Big Sink Unit, the upper stratigraphic member of the Phosphate Mine Beds (Osborne, 1983). It consists of carbonate-cemented osseous sandstones interbedded with thin layers of structureless mud (Osborne, 1983). The unit is hypothesised to be older than (i.e. stratigraphically below) the breccia of the nearby bone-rich Bone Cave, identified by Osborne (1983) as belonging to the Mitchell Cave Beds. The Bone Cave fauna contains many species typical of Pleistocene deposits of eastern Australia (Dawson, 1985). Correlation of associated fauna with taxa otherwise known from better-dated deposits in Queensland and Victoria suggests that the Big Sink deposit from which the holotype was obtained is probably early Pliocene in age.

Description (based on the holotype and paratypes). Because the new species is clearly the closest sister­group of Macroderma gigas (see Discussion), it is described only in so far as it differs from M. gigas.

Maxillary fragments of M. koppa preserve cheek and orbital regions and part of nasal region; no sutures visible. In contrast to M. gigas, with single infraorbital foramen, 2 infraorbital foramina exhibited by this species (Fig. 3). Larger, anterior foramen opens onto face within fossa developed above anterior end of P4. Second, smaller foramen penetrates maxilla, possibly for lateral branch of trigeminal (facial) nerve, occurs within fossa developed above posterior end of P4. Larger fossa directed more anteriorly towards what would have been noseleaf; smaller fossa is directed anteroventrally towards lips or cheeks.

Nasal region of maxilla rises at approximately same angle as that of M. gigas (i.e. about 45 0 with respect to tooth row), is similarly gently concave. However, at least anterior part of nose-shield differs markedly from M. gigas in lack of bony lateral eminences (Fig. 4). Posterior (dorsal) region of nose­shield not preserved.

In their teeth, there is a great deal of morphological overlap between the two species. However, a combination of features in the anterior dentition distinguishes M. koppa from M. gigas. These features of M. koppa are described below.

Cl, represented by 6 specimens, semicircular in basal outline with buccal side convex. Base of tooth posterobuccally swollen with marked notch separating base of paracone from base of large posterolingual accessory cusp. Latter cusp appears to be more discrete or isolated in M. koppa than M. gigas, being smaller in cross-sectional area and rising more steeply from base of crown. Posterolingually, tall but narrow cingUlum commonly surrounds entire tooth but does not always enclose posterolingual accessory cusp. Cl of M. koppa is comparable in size to Cl oflarge specimens of M. gigas but lacks marked anteroposterior compressional flexure in base of lingual cingulum which characterises M. gigas.

p4 is represented by 5 specimens which are comparable in width to, but much longer than, those of M. gigas. Difference in length appears due to elongation anterior to paracone so that principal cusp situated far more posteriorly along buccal edge of tooth than in M. gigas, accentuated by posteriorly recurved shape of paracone in M. koppa, shallow­angled rise of anterior crest of paracone from anterior cingUlum and overall lower crown height of tooth. Anterior cingulum more pronounced in M. koppa with a cingular swelling occurring at its highest point. In lingual cingulum of broad heel, a distinct cingular cusp is developed which is also variably present within populations of M. gigas.

The morphology of Ml, M2 and M3 appears to fall within the range of variation observed in M. gigas.

Lower incisors of M. koppa, known from holotype and UNSWWC156, farmorerobustthan thoseofM. gigas, being at least one and one half times larger in crown area. Of what is known of incisors, their shape aparently similar to those of M. gigas with lateral incisors being much larger than inner incisors.

Cl' represented by 6 specimens, similar in both size and shape to those of large individuals of M. gigas. Some intra specific variation in M. koppa, however, appears to occur at lowest posterobuccal point of tooth (i.e. base of buccal edge ofposteriorly flattened face of protoconid) where there is sometimes a break in otherwise high buccal cingUlum.

P 2' represented by 3 specimens, show considerable variation in depth and height of buccal cingUlum. Tooth much wider and longer than in M. gigas, being markedly longer posterior to protoconid, so that in M. koppa protoconid occurs approximately midway along length of tooth.

P 4' represented by 5 specimens, noticeably more robust than in M. gigas (Hand, 1985, fig. 12), being longer, wider, markedly less laterally compressed in area of protoconid. Longitudinal crest linking tip of protoconid to point on posterior cingulum appears to

Hand, Dawson & Augee: Macroderma koppa 347

MM • • • • •

Fig.3. Comparison of the maxillas and number of infraorbital foramina in: a-a! (stereopairs), Macroderma koppa n.sp. (UNSWWC171); and b-b! (stereopairs), Macroderma gigas (WAM 65.12.191). Photos by R. Oldfield.

be more pronounced in M koppa, often changes angle at base of protoconid. On either or both sides of crest (i.e. posterolingually and/or posterobuccally) depressions may be developed.

Morphology of Mp M2 and M3 (represented by minimum of7 specimens each) appears to fall within range of variation in morphology observed in M. gigas (Hand, 1985, figs 13-15). In M! and M2,

however, buccal cingulum swollen and ornate at point just anterior to protoconid. This feature, which occurs variably in populations of M. gigas (e.g. Hand, 1985, fig. 14), appears to be fixed in this population of M. koppa (but see below).

Table 1 gives measurements of the holotype, paratypes and refered specimens made to the nearest O.lmm using a Wild MMS235 Digital Length-

348 Records of the Australian Museum 40

MM • • • • • Fig.4. Comparison of the development of the bones comprising the frontal nose-shield in: a-a l

(stereopairs), Macroderma koppa n.sp. (UNSW WCl7l); and, b-b l (stereopairs), Macroderma gigas AM MlOl90). Photos by R. Oldfield.

Measuring Set attached to a Wild M5A Stereomicroscope (see Hand, 1985, figs 9-10 for further explanation of measures).

Further comparisons. Other large specimens of Macroderma examined in this study include a dentary fragment from Australian Museum fossil collections (AM F4 7021; see Molnar et aI., 1984, fig. 3) collected from an unknown locality in the

Wellington Caves system. The specimen was probably collected last century or early this century and originally housed in the Mining Museum, Sydney before transfer to the Australian Museum (see Dawson, 1985).

Features of AM F47021 shared with M. koppa include its large size, lack oflateral compression ofP 4

and its pronounced crest on P4 running from the tip

of the protoconid to the posterior cingulum. For these reasons, the dentary fragment AM F47021 is tentatively referred to M. koppa n.sp. This specimen does not, however, exhibit a particularly swollen or ornate buccal cingulum on M I_2•

Another specimen from "Wellington Caves", BMNH 42670b, discussed by Molnar et al. (1984) and housed in the British Museum, was not examined in this study.

Large Quaternary specimens of Macroderma from Yessabah Caves in north-eastern New South Wales and Cliefden Caves in eastern central New South Wales (see Fig. 1) are currently refered to M. gigas (Molnar et al., 1984). Although larger than most specimens of M. gigas, because they are represented only by isolated dentaries lacking anterior teeth their identity remains in doubt.

The maxilla of a recently described partial skull recovered from Ashford Cave, north-eastern New

Hand, Dawson & Augee: Macroderma koppa 349

South Wales (see Fig. 1) (Macroderma sp. ofMolnar et al., 1984, figs 1,2) exhibits a single infraorbital foramen which excludes it from M. koppa. The lingual shape of the M3 protocone, the feature singled out by Molnar et al. as being different, does not appear to differ significantly from larger samples of M. gigas and it is therefore refered to that species.

Etymology. The species is named after "Koppa", the mythological spirit believed by Australian Aborigines to have inhabited Wellington Caves (Lane & Richards, 1963). The reference to a spirit or ghost alludes to this species' close phylogenetic relationship with the living Ghost Bat, Macroderma gigas. Of the Big Sink fossil species so far identified, M. koppa, like its mythological counterpart, is probably the only one that actually inhabited the cave.

Table 1. Measurements of upper and lower teeth of the holotype (AM F69070), paratypes and referred specimens of Macroderma koppa n.sp. from Wellington Caves, New South Wales. In mm. (See Hand, 1985, figs 9-10 for further explanation of measures.)

Character

Length CI_M3 Length p4_M3 Length MI_M3 p4 buccal length M I buccal length M2 buccal length M3 buccal length p4 lingual length M I lingual length M2 lingual length p4 width MI width M2width M3 width Dentary depth below

M2 protocomid Length CI-M3 LengthPc M3 Length M I-M3 P41ength P 4 anterior cingulum

to protoconid length

Mllength (in situ) M2length (in situ) M31ength (in situ) M I trigonid length M I talonid length M2 trigonid length M2 talonid length M3 trigonid length M3 talonid length P4 width MI trigonid width M I talonid width M2 trigonid width M2 talonid width M3 trigonid width M3 talonid width

Holotype Paratypes and referred specimens AM UNSW UNSW UNSW UNSW UNSW UNSW UNSW UNSW UNSW UNSW UNSW AM

F69070 WCl52 WCl62 WCl61 WC163 WC154 WC158 WC153 WC164 WC156 WC159 WCI70 F47021 16.7 13.1 12.8 9.4 4.1 4.2 3.7 4.0 4.1 4.1 4.0 4.2 3.8 3.6 3.8 3.9 4.2 2.0 1.9 4.1 4.2 4.0 3.6 3.9 3.6 3.5 3.2 4.4 4.3 4.1 4.5 4.4 4.5 4.6 4.1 4.3 4.5

4.2 4.5 19.7 15.0 12.1 3.4 3.2

1.4 4.2 3.7 4.2 4.1 4.0 2.7 2.3 1.5 1.5 2.5 2.4 1.6 1.6 2.5 1.4 2.3 2.4 2.3 2.5 2.4 2.8 2.4 2.6 2.3 2.5 2.1

2.5 1.4

2.0 2.3 2.3

2.2 1.6 2.3 1.3

2.1 2.2 2.4 2.2

2.5 1.4

2.2 1.6

4.4

3.1

1.4 3.9 4.0

2.6 1.4

2.3 2.5 1.6 1.5 2.2 2.5 1.7

2.1 2.4 2.5

2.4 2.6 2.3 2.5 2.4 1.9

3.6

1.9 3.8 3.9

2.5 1.4 2.4 1.5

2.1 2.2 2.4 2.5 2.5 2.6

350 Records of the Australian Museum 40

Discussion Recent geological studies by Frank (1971, 1972,

1975), Francis (1973) and Osborne (1983) have shown that bone-bearing stratigraphic units in Wellington Caves are not contemporaneous. Osborne (1983) suggests that they represent a minimum of three periods of deposition ranging from at least the Pliocene to late Pleistocene. The Big Sink Unit has been identified as being the upper member of the Phosphate Mine Beds (Osborne, 1983) and, on the basis of faunal evidence, is hypothesised to have been deposited during the early to middle Pliocene.

This faunal evidence is provided by a number of studies in progress involving stage-of-evolution comparisons with mammalian taxa from the stratigraphically higher (i.e. younger) Bone Cave sediments, as well as other better-dated Australian Pliocene and Pleistocene fossil mammal assemblages. For example, the dominant macropodine of the Big Sink local fauna, Protemnodon sp. cf. P. devisi, differs only slightly

(Dawson, 1985) from P. devisi which is known only from the early Pliocene-aged Chinchilla Sand fauna of south-eastern Queensland (Bartholomai, 1973). A Pliocene age is also supported by the presence of Thylaeoleo erassidentatus and, among the small marsupials, Petauroides stirtoni and a unique peramelid which are otherwise known only from the early Pliocene Hamilton local fauna of Victoria. A Pliocene age is also indicated by the conspicuous absence of many species which are typically found in Pleistocene deposits (e.g. species of Maeropus, Proeoptodon, Diprotodon optatum, Sthenurus atlas, several rodent and dasyurid species, etc.) and which are abundantly present in the Bone Cave deposits of Wellington Caves (Dawson, 1985).

The Big Sink's Macroderma koppa is clearly distinguished from all known modern and Pleistocene specimens of M. gigas by the retention of two infraorbital foramina on the facial side of the maxilla. The double foramen state is a plesiomorphic (or primitive) condition present in most megadermatids (Hand, 1985) with the exception of

Fig.5. A cladogram of the hypothetical relationships of Australian megadermatids (where nycterid and other rhinolophoid species have been used to determine polarity of character-states within the Megadermatidae; see text and Hand, 1985). Potential apomorphies are as follows: 1, robust forms, enlarged heel of P4_M2, paracone of Cl relatively more prominent, increase in the robusticity of the buccal cingula of lower molars; 2, (see systematic diagnosis and description in Hand, 1985; but autapomorphies are unclear); 3, increase in size, increase in relative crown height, convergence of principle cusps and crests on each crown, reduction ofpreentocristids, loss of p2, posterolingual expansion of the heel of P4, M2 hypoconulid more posteriorly-situated and better­developed, p4 parastyle better-developed; 4, increase in size, pronounced lingual cingular cusp on P4, more robust buccal cingula on lower molars; 5, single infraorbital foramen, relative shortening of the anterior tooth row, elaboration of frontal nose-shield, lateral compression of P 4' higher crowned and less posteriorly recurved P4.

species of the French early Tertiary genus Necromantis (Revilliod, 1922) and all specimens of M. gigas. Necromantis adichaster and M. gigas exhibit a single infraorbital foramen. Because N. adichaster lacks most of the derived features that M. gigas shares with other living and. fossil megadermatids (e.g. loss ofP J' a posteriorly extended palate, well-developed heels on P4_M2 and an expanded paroccipital shelf) and exhibits a number of striking autapomorphies (e.g. a conspicuously flattened skull, shortened face and inclined rectilinear sagittal crest), the shared state of the single foramen in N. adichaster and M. gigas is interpreted to be the result of evolutionary convergence (Hand, 1985).

M acroderma koppa and M. gigas appear to be very closely related. M acroderma gigas appears to be a relatively recent offshoot from an ancestral population of the kind represented by M. koppa. A phylogenetic hypothesis involving these species and the Australian Miocene M. godthelpi is given in Fig. 5. Determination of the polarity of character-states in this phylogenetic analysis has been facilitated by common recognition (following Miller, 1907) of a Megadermatidae-Nycteridae sister-group relation­ship within the Rhinolophoidea (see Hand, 1985).

Features that distinguish M. koppa from M. gigas include its relatively poorly-developed nose-shield, more robust incisors and premolars, less laterally compressed P 4' lower crowned but larger p4 and its proportionately longer anterior tooth row. In these plesiomorphic features, M. koppa more closely resembles the Australian Miocene species Macroderma godthelpi than the living M. gigas (Fig. 5).

M acroderma koppa is the second fossil megadermatid to be described from Australia. Four new but as yet undescribed Miocene megadermatids and a tiny Pliocene species have been recently recovered from deposits on Riversleigh Station (Hand, 1987). This diversity indicates that M. godthelpi, M. koppa and M. gigas are representatives of what was once an extensive Australian radiation of false vampires, second only, perhaps, to that of the European Tertiary.

ACKNOWLEDGEMENTS. This research was funded by grant number A3 83/15255 from the Australian Research Grants Scheme to M. Augee. During this study, S. Hand was supported by a Commonwealth Postgraduate Research Award and a Department of the Arts, Sport, the Environ­ment, Tourism and Territories Research Fellowship.

We thank Mr R. Jones of the Australian Museum for providing access to fossil specimens vital for comparisons with the new species; Dr D. Kitchener (Western Australian Museum) and Ms L. Gibson (Australian Museum) for kindly lending modern Macroderma specimens for comparative purposes; Dr R. Oldfield for producing the photographs; and Ms J. Taylor for helping to produce the other figures.

Constructive criticism given by Dr T. Flannery and an anonymous referee is gratefully acknowledged.

Hand, Dawson & Augee: Macroderma koppa 351

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Accepted 12 September 1988